Host selection byHylemya antiqua (Meigen)

Volatile components ofAllium cepa L, trapped in and recovered from Porapak Q, were separated by gas-liquid and thin-layer chromatography. Fractions eliciting oviposition fromHylemya antiqua (Meigen) were analyzed by combined gas chromatography-mass spectroscopy (GC-MS) and were found to contain saturated and unsaturated disulfides. Authentic samples of the identified components were obtained by synthesis and tested for oviposition response byH.antiqua. Three new attractants and oviposition stimulants forH. antiqua are methyl propyl disulfide,cis-propenyl propyl disulfide, andtrans-propenyl propyl disulfide. All were present in greater proportion in the volatiles from fresh onions than from stored onions. Dimethyl disulfide, methylcis-propenyl disulfide, and methyltrans-propenyl disulfide did not elicit significant oviposition fromH. antiqua.Diptera: Anthomyiidae.Research supported by National Research Council of Canada, Operating Grant Nos. A3881 and A3706, and Cooperative Grant No. A0243: Nature Products Chemistry and Chemical Ecology of Canadian Insect and Disease Pests.     

Aggregated oviposition inDelia antiqua (Meigen): A case for mediation by semiochemicals

Experiments conducted in the laboratory tested the hypotheses that aggregated oviposition by onion maggot flies,Delia antiqua (Meigen), is caused by stimuli associated with ovipositing females, newly laid eggs, or both. Using a paired oviposition station bioassay that eliminated visual stimuli associated with the treatment under study, 67% of the eggs laid by caged females were in response to the odor of females already ovipositing on an onion slice, as opposed to 33% of the eggs laid in response to an onion slice alone. When newly laid eggs were transferred to onion slices and held for either 24 or 48 hr before being bioassayed against similarly aged untreated onions, 74% and 97% of the eggs were laid at the egg-treated onion stations, respectively. Similar results were achieved when an aqueous wash of newly laid eggs was applied to the onion slice. When the egg wash was processed through a bacterial filter or when eggs were present but not in contact with onions, all response was eliminated. These results implicate microorganisms transmitted on the egg surface in creating an attraction for ovipositing females. Heptane extracts of ovipositor tips from mated, ovipositing females induced 72% of the test females to oviposit near points at which extracts were applied to the oviposition station floor. A behavioral sequence for an optimal host-selection strategy is hypothesized, whereby host-seeking female onion flies respond to host-derived alkyl sulfides at long range and metabolic by-products of microbially infested hosts and visual cues at short range (ca. l m), with final selection of oviposition sites potentially reinforced by contact with an aggregation pheromone released or left on the substrate by ovipositing females.     

Production of onion fly attractants and ovipositional stimulants by bacterial isolates cultured on onion

Decomposing onions at certain microbial successional stages produce potent volatile attractants and ovipositional stimulants of the onion fly,Delia antiqua (Diptera: Anthomyiidae). A reproducible source of these compounds was obtained by culturingErwinia carotovora var.carotovora (EC) on sterile onion tissue. In laboratory choice tests, EC-inoculated onion was more attractive thanKlebsiella pneumoniae (KP) cultured on onion, EC cultured on potato (a nonhost of onion fly), or the chemical synthetic baits dipropyl disulfide and an aqueous solution of 2-phenylethanol and pentanoic acid. Onion flies were mildly attracted to potato after inoculation with EC, but females did not accept EC-inoculated potato for oviposition. This work emphasizes that sources of semiochemicals may need to be defined microbiologically as well as physically and chemically.Diptera: Anthomyiidae.Paper No. 12106 of the Michigan State University Agricultural Experiment Station.     

Suppression of oviposition inOryzaephilus surinamensis (L.) (Coleoptera: Cucujidae) following prolonged retention in high-density cultures or short-term exposure to larval volatiles

After 20 days in a high-density culture containing many larvae, female sawtoothed grain beetles,Oryzaephilus surinamensis (L.), laid half as many eggs in a 24-hr oviposition bioassay as females held for six days in the same culture, or for six or 20 days in a low-density culture. Oviposition by females held for six days in a high-density culture was reduced to a similar extent when they were exposed in the oviposition bioassay to an oat flake treated with an extract of Porapak Q-captured larval volatiles (equivalent to 5000 larval hours). A retained suppression of oviposition rate after prolonged exposure to larvae or an induced reduction caused by short-term exposure to larval volatiles both could be of adaptive advantage in reducing the risk of oviposition in an already densely populated habitat.Research supported by the Natural Sciences and Engineering Research Council of Canada, Strategic Grant G0958 and Operating Grants A3881 and A3785.     

Sex pheromone components and calling behavior of the female dermestid beetle,Trogoderma variabile Ballion (Coleoptera: Dermestidae)

Females of the dermestid beetle,Trogoderma variabile Ballion, exhibit a diel pattern of calling behavior. A potent sexual excitant and attractant for the male beetles can be collected on Porapak Q during aeration of female beetles, but cannot be detected in extracts of macerated females. The attractant has been identified as (Z)-14-methyl-8-hexadecenal. Of the five additional compounds previously identified as attractants in otherTrogoderma species, only two were found; (Z)-14-methyl-8-hexadecen-1-ol is present in extracts of macerated female beetles, but not in extracts of Porapak Q, and -caprolactone is present in extracts of Porapak Q. Three collection procedures were necessary to ensure that all the pheromone components had been isolated. Synthetic (Z)-14-methyl-8-hexadecenal elicits attraction and sexual arousal inT. Variabile males.Mention of a commercial or proprietary product in this paper does not constitute an endorsement of that product by the USDA.     

Influence of cage design on precision of tube-trap bioassay for attractants of the onion fly,Delia antiqua

Responses of onion flies,Delia antiqua, to known attractants were measured in the laboratory with a novel tube-trap bioassay. The relative numbers of flies caught in tube traps baited with enzymatic yeast hydrolysate, brewer's yeast, andn-dipropyl disulfide were similar to those obtained previously with cone traps in the field. Changing the shape of the bioassay cage from a cuboid to a cylinder decreased the experimental error obtained from analysis of variance, as did rotating the floor of the circular cage. This bioassay should be useful in evaluating attractants for other insects that orient along the substrate.Diptera: Anthomyiidae.Paper No. 11327 of the Michigan State University Agricultural Experiment Station. Received for publication June 8, 1984.     

Cherry Bark Tortrix, Enarmonia formosana: Olfactory Recognition of and Behavioral Deterrence by Nonhost Angio- and Gymnosperm Volatiles

We tested the hypothesis that males and females of the cherry bark tortrix (CBT), Enarmonia formosana, antennally perceive and behaviorally respond to volatiles from nonhost plants. Volatiles from flowering cherry trees, Prunus serrulata Kwanzan, and from nonhost trees, including trembling aspen, Populus tremuloides, grand fir, Abies grandis, Norway spruce, Picea abies, and Scots pine, Pinus sylvestris, were captured on Porapak Q and extracts analyzed by coupled gas chromatographic–electroantennographic detection (GC-EAD). Compounds that elicited responses from male and female antennae were identified by coupled GC–mass spectrometry (MS) and by comparative GC-MS and GC-EAD with authentic standards. In field cage and field experiments, nonanal from aspen trees and (±)--pinene from coniferous trees were effective in reducing captures of male CBT in pheromone-baited traps and deterring oviposition by female CBT on cherry blocks. Recognition of nonhost volatiles, such as nonanal and -pinene, may allow male and/or female CBT to avoid trees that provide inadequate concealment from predators during calling, resting, and copulation and that are unsuitable for oviposition and development of offspring. Nonhost volatiles may also be exploited for control of CBT by disrupting both mate- and host-finding.     

Onion fly and little house fly host finding selectively mediated by decomposing onion and microbial volatiles

Responses of onion flies,Hylemya antiqua (Meigen), to various synthetic onion and microbial volatiles as well as volatiles from microbial cultures and decomposing onions were tested to characterize the most effective host-finding stimuli. Of nine onion and microbial volatiles tested individually, only the known attractant,n-dipropyl disulfide, caught significant numbers of flies. However, a blend of these volatiles attracted more flies than any single chemical, includingn-dipropyl disulfide. In another experiment, agar plates inoculated with microorganisms from decomposing onions did not attract onion flies. However, cut onions inoculated with microorganisms and conditioned 4 days caught more onion flies than freshly cut onions andn-dipropyl disulfide. These results suggest that a blend of chemicals, rather than a single key chemical, is the more effective host-finding stimulus, and that microbial activity enhances the attractancy of a blend of onion volatiles. Large numbers ofFannia canicularis (L.), the little house fly, responded to the microbial cultures, demonstrating the existence of a potent attractant for this muscid.Diptera: Anthomyiidae.Diptera: Muscidae.Paper No. 9470 of the Michigan State University Agricultural Experiment Station. Received for publication May 23, 1980.     

Monoterpene oviposition stimulants ofDioryctria amatella in volatiles from fusiform rust galls and second-year loblolly pine cones

Volatiles collected on Poropak Q from fusiform rust galls (Cronartium quercuum F. sp.fusiforme) of loblolly pine for 3 hr were better ovi-position stimulants forDioryctria amatella (Hulst) females than extracts of 8-hr collections. GLC analysis of these extracts showed no major differences in relative monoterpene composition, although 8-hr collections contained an unidentified compound not detected in the 3-hr collections. Comparison of volatiles from second-year loblolly pine cones with those from fusiform galls showed that both containeddl--pinene, (–)--pinene, myrcene, and (+)-limonene. Camphene was found in galls only, and relatively large quantities of -phellandrene were identified only from cones. The five major monoterpenes found in the two host substrates were tested in an oviposition bioassay. Eighteen trials using different combinations of these terpenes showed that the combination of -pinene, myrcene, and limonene was as attractive toD. amatella females as all other terpene combinations, including turpentine.Lepidoptera: Pyralidae.     

Lanierone: A new pheromone component fromIps pini (Coleoptera: Scolytidae) in New York

A new pheromone component, lanierone, (2-hydroxy-4,4,6-trimethyl-2,5-cyclohexadien-1-one) was isolated and identified from a Porapak Q collection of volatiles from maleIps pini from New York through GC fractionation, bioassay, and spectrometry. In both the laboratory and the field, synthetic lanierone, in a 1:100 ratio with synthetic ipsdienol, is as attractive as natural pheromone sources. Synthetic ipsdienol alone is not attractive in the laboratory and only weakly attractive in the field. Varying the ratio of lanierone to ipsdienol in the field from 10^–41 to 11 in 10-fold increments resulted in an increased number of beetles trapped at the three lower ratios, but also in an increase in the proportion of males trapped. In the field, all combinations of lanierone to ipsdienol attracted proportionately fewer males than did pheromone-producing male beetles. GC and GC-MS analyses of Porapak Q-trapped volatiles revealed that lanierone is produced in an amount equal to about 0.2% of that of ipsdienol and is produced exclusively by males. The small amount of lanierone produced, together with a GC retention time similar to that of ipsdienol on a nonpolar column, probably confounded its detection in earlier studies.Deceased.     

Components of moribund American elm trees as attractants to elm bark beetles,Hylurgopinus rufipes andScolytus multistriatus

Hylurgopinus rufipes male and female beetles were attracted to American elms infected with Dutch elm disease, and to American elms killed by injection of cacodylic acid.H. rufipes was also attracted to solvent extracts of elm, or to Porapak Q-trapped volatiles from elm. The major components of attractive fractions of Porapak Q-trapped volatiles were isolated, identified, and tested in field bioassays. Several artificially compounded mixtures of sesquiterpenes were attractive toH. rufipes, although no bait tested was as attractive as diseased tree controls. Laboratory bioassays withH. rufipes were marginally successful. In laboratory bioassays, nine of 14 sequiterpenes identified from active fractions of Porapak extracts elicited significant response fromScolytus multistriatus male and female beetles: - and -cadinene, -cubebene, -muurolene, and -elemene were most active. However, in field tests, none of the sesquiterpenes alone or in combination significantly attractedS. multistriatus, nor did they significantly enhance the attraction ofS. multistriatus to female-produced pheromone components (4-methyl-3-heptanol [H] and -multistriatin [M]). In other field tests, -cubebene (C) significantly enhanced response ofS. multistriatus to H plus M, but foliage, logs, or chips of healthy elm did not enhance trap catch to HMC.Coleoptera: Scolytidae.     

Factors Accounting for the Variability in the Behavioral Response of the Onion Fly (Delia antiqua) to n-Dipropyl Disulfide

The onion fly Delia antiqua is a specialist herbivore attacking only onions and closely related Allium species. n-Dipropyl disulfide (Pr₂S₂) has long been known to be attractive to onion flies in the laboratory and in the field. However, the insect's response is highly variable. Using behavioral bioassays we found that Pr₂S₂ was highly attractive to gravid, mated female onion flies, but did not stimulate oviposition. The response of female onion flies was concentration dependent. The physiological state of the flies (i.e., age, mating status, egg load/oviposition experience) also affected their responsiveness. The response of both sexes of onion flies varied with age, but females were always more strongly attracted than males. Responsiveness of females increased during the first 10 days after emergence. It stayed at a high level until 21 days after which the experiment was terminated. The responsiveness of males reached a maximum at 6–7 days after which it declined. Mated, gravid females responded more strongly to Pr₂S₂ than unmated, gravid females. Females deprived of the opportunity to oviposit were more attracted compared to females that had oviposited on cut onions prior to the experiment. Electroantennograms (EAG) of females revealed a higher response to stimulation compared to males. The EAG-response of females was not affected by mating status.     

Dodecanoic Acid Is the Oviposition Pheromone ofLutzomyia longipalpis

The sandfly,Lutzomyia longipalpis (Lutz and Neiva), produces an oviposition pheromone in the accessory glands that is secreted onto the eggs during oviposition. This compound attracts and/or stimulates gravid females for egg-laying. The compound was identified as dodecanoic acid, using gas chromatography–mass spectrometry, gas chromatography, and chemical derivatizations. The synthetic analog induced the same behavioral response in gravid sandflies as the whole egg extract when present in biologically relevant quantities. When single sensillum recordings were taken from the ascoid on the antennae of female sandflies, the response to egg extract and dodecanoic acid was similar. There was a dose dependent response to dodecanoic acid and preferential sensitivity compared to other fatty acids. There was a strong additive interaction upon the behavior ofL. longipalpis when dodecanoic acid was tested in a bioassay with the known oviposition attractant apneumones from rabbit feces, hexanal, and 2-methyl-2-butanol. The results suggested that sandflies acquired hexadecanoic acid (palmitic acid) from the blood meal and over a period of four days this was converted to dodecanoic acid. The role of these token semiochemicals in sandfly oviposition is discussed with respect to additive interactions and the significance of their origin from larval food resources.     

Aggregation pheromone of the male granary weevil,Sitophilus granarius (L.)

Evidence for a male-produced aggregation pheromone inSitophilus granarius is reported. Hexane extracts of Tenax®-trapped volatiles from males held on wheat were attractive to both sexes in a multiple-choice olfactometer and pitfall bioassay chamber. A quantitative relationship existed between days of insect exposure on disks and degree of responsiveness. The maximum response was to 35 insect-day-equivalents. Diel-related activity showed both sexes responsive during photophase and nonresponsive during scotophase.Coleoptera: Curculionidae.Contribution to Regional Research Project NC151, Marketing and Delivery of Quality Cereals and Oilseeds in Domestic and Foreign Markets.Mention of a commercial or proprietary product does not constitute an endorsement by the USDA.     

1-Octen-3-ol,attractive semiochemical for foreign grain beetle,Ahasverus advena (Waltl) (Coleoptera: Cucujidae)

Volatiles were captured on Porapak Q from foreign grain beetles,Ahasverus advena (Waltl), feeding on rolled oats at various population densities. At low population density, males, females, and mixed-sex beetles four to six weeks posteclosion and older produced 1-octen-3-ol. Mixed-sex beetles emitted almost pure (R)-(–) enantiomer. Weekly production rates of 1-octen-3-ol by males were at least four times greater than those of females. Production of 1-octen-3-ol was barely detectable in volatiles from mixed-sex adults maintained at the highest population density. Laboratory bioassays in a two-choice, pitfall olfactometer modified to retain responding beetles revealed that 1-octen-3-ol serves as an aggregation pheromone forA. advena. Both racemic and chiral 1-octen-3-ols were good attractants for mixed-sex adults in the pitfall olfactometer.Research supported by the Natural Sciences and Engineering Research Council of Canada, Strategic Grant G0958 and Operating Grants A3881 and A3785.     

Induction and Systemic Release of Herbivore-Induced Plant Volatiles Mediating In-Flight Orientation of Aphidius ervi

In-flight orientation of the braconid Aphidius ervi in response to volatiles released from broad bean plants infested by the pea aphid, Acyrthosiphon pisum, was studied in a no-choice wind-tunnel bioassay. The role of aphid infestation level and duration, systemic production of volatiles by insect-free parts of the plant, and the specificity of aphid-induced volatiles on the flight behavior of the foraging female parasitoids were investigated. The upper insect-free part of a three-leaved broad bean plant, which was basally infested by a population of 40 A. pisum, released synomones detectable by A. ervi females after at least 48–72 hr of infestation, resulting in both significant increases in oriented flights and landings on the source compared with uninfested control plants. This suggests that volatiles involved in host-location by A. ervi are systemically released by broad bean plants either in response to circulation of aphid saliva, circulation of saliva-induced bioactive elicitors, or circulation of the synomones themselves. Air entrainment extracts of volatiles collected from a broad bean plant infested by the nonhost Aphis fabae or an uninfested broad bean plant elicited few oriented flights and landing responses by female parasitoids. These extracts were significantly less attractive than extracts collected from a broad bean plant infested by the host A. pisum, indicating the specificity of synomones elicited by different aphid species on the same plant species.     

Primary Attraction of the Fir Engraver, Scolytus ventralis

In laboratory bioassays, Porapak Q-captured and steam-distilled volatiles from the bark of host trees, Abies grandis, particularly from root-rot-infected trees, attracted 50–70% of male and female fir engravers, Scolytus ventralis. Gas chromatographic–electroantennographic detection (GC-EAD) analyses of Porapak Q-captured bark volatiles revealed 19 EAD-active compounds of which 13 (mostly monoterpenes) were identified by GC–mass spectrometry (GC-MS). In separate field experiments, multiple-funnel traps baited with two blends of these 13 synthetic volatiles released at 280 and 340 mg/ 24 hr attracted 66 and 93% of the total S. ventralis captured, respectively. The clerid predator, Thanasimus undulatus, also responded strongly to the kairomonal volatiles. Additional experiments produced no evidence for aggregation pheromones in S. ventralis. These included laboratory bioassays and GC and GC-EAD analyses of Porapak Q-captured volatiles from male- and female-infested logs or trees undergoing mass attack in the field, GC analyses and/or bioassays of extracts from female accessory glands, extracted volatiles from emerged, attacking and juvenile hormone-treated beetles of both sexes, and videotape analysis of the behavior of attacking beetles on the bark surface. We argue against the hypothesis of pheromone-mediated secondary attraction in S. ventralis and conclude that the attack dynamics of this species can be explained solely by its sensitive primary attraction response to host volatiles.     

A quantitative feeding bioassay forPissodes strobi Peck (Coleoptera: Curculionidae)

A feeding bioassay forPissodes strobi Peck was developed that consisted of plastic petri dishes containing paired agar disks immersed in paraffin wax. Lens paper covering the top surface of the disks provided 1, 2, or 3 weevils in each replicate with a surface through which they made regular feeding punctures which could be easily counted. The number of feeding punctures was correlated with the amount of agar ingested. Candidate feeding stimulants or deterrents were applied to the paper covering one of the disks in the dishes, while the other served as a solvent control. Feeding stimulants were tested using disks of pure agar, and deterrents were assayed on agar disks that contained 2% ground, dried Sitka spruce bark,Picea sitchensis Carr. The weevils exhibited a concentration-dependent response to the amount of spruce bark in the agar disk. Feeding stimulation by Sitka spruce bark extracts, and deterrency by leaf oil of western red cedar,Thuja plicata Donn, was demonstrated. The bioassay would be useful in chemical isolation programs aimed at identification of feeding stimulants and deterrents.Research supported by National Research Council of Canada Cooperative Grant A0243 and Operating Grants A3881 and A3706.     

3-methyl-3-buten-1-ol: An aggregation pheromone of the four-eyed spruce bark beetle,Polygraphus rufipennis (Kirby) (Coleoptera: Scolytidae)

Porapak Q-captured volatiles of malePolygraphus rufipennis (Kirby) boring in black spruce and white spruce from Newfoundland and British Columbia, respectively, contained 3-methyl-3-buten-1-ol. Volatiles from logs in which the males had been joined by females contained the compound in reduced quantities. Hindgut extracts from male and femaleP. rufipennis disclosed no male-specific volatiles, but 3-methyl-3-buten-1-ol was detected in extracts of male-produced frass. The compound was not present in extracts from fresh phloem tissue.P. rufipennis of both sexes responded strongly in the field to traps baited with 3-methyl-3-buten-1-ol released at 4390 g/day. There was little response to unbaited traps, fresh uninfested black spruce logs, or to 3-methyl-3-buten-1-ol released at lower rates. Combination of 3-methyl-3-buten-1-ol with either one of five terpenes prevalent in black spruce did not enhance beetle catch. Approximately half of 20 black spruce trees baited with 3-methyl-3-buten-1-ol were attacked, compared to 20.0% of 10 unbaited control trees. This new aggregation pheromone could be used to monitor or manageP. rufipennis populations.     

Pheromones in white pine cone beetle,Conophthorus coniperda (schwarz) (Coleoptera: Scolytidae)

Female white pine cone beetles,Conophthorus coniperda, attacking second-year cones of eastern white pine,Pinus strobus L., produced a sex-specific pheromone that attracted conspecific males in laboratory bioassays and to field traps. Beetle response was enhanced by host monoterpenes. The female-produced compound was identified in volatiles collected on Porapak Q and in hindgut extracts as (+)-trans-pityol, (2R,5S)-(+)-2-(1-hydroxy-1-methylethyl)-5-methyltetrahydrofuran. Males and females produced and released the (E)-(-)-spiroacetal, (5S,7S)-(-)-7-methyl-1,6-dioxaspiro[4.5]decane, which was not an attractant for either sex, but acted as a repellent for males. Porapak Q-trapped volatiles from both sexes contained (+)-trans-pinocarveol and (-)-myrtenol. In addition, hindgut extracts of females containedtrans-verbenol, while males had pinocarvone and verbenone. Work in Georgia and Canada confirmed that the same isomers of pityol and spiroacetal are present in two distinct and widely separated populations ofC. coniperda.